Why does onlyAthalia Japonica enter summer diapause among three sympatricAthalia sawflies feeding on crucifers?

Summary This report assesses the primary factor for the evolution of summer diapause of the three species of sawfly,Athalia japonica, A. rosae andA. infumata that feed on cruciferous plants and coexist in the same area.A. japonica has two discrete spring and autumn generations, butA. rosae andA. infumata 5–6 generations. OnlyA. japonica enters summer diapause in response to the long daylengths in spring. Although these three sawflies usually feed on the same cultivated crucifers, they differ markedly in the utilization of wild crucifers. They oviposit only on young leaves.A. japonica mainly usesCardamine plants which sprout in spring and autumn.A. rosae andA. infumata primarily use hosts with new leaves all the year round, i.e. cultivated crucifers andRorippa indica, respectively. The thermal threshold for development is lower inA. japonica than in the other two species. The low heat tolerance ofA. japonica is adapted only to cool shady habitats whereCardamine grows. Presumably, summer diapause ofA. japonica is adaptation to the deterioration of the primary host plants rather than unfavorable climatic conditions. This interpretation is supported by the movement patterns of the threeAthalia sawflies, alternative means to escape from deteriorated habitat conditions.     

Movement patterns of threeAthalia sawflies in relation to the spatio-temporal distributions of their habitats

Summary I compared the differences in the movement intensity of three species of sympatricAthalia sawflies,A. japonica, A. rosae andA. infumata feeding on cruciferous plants. Mark-release-recapture census was conducted to estimate movement distance, sex ratio and age composition of adult sawflies. In addition, the sex ratio of newlyemerged adults was examined by rearing field-collected larvae until adult emergence. Age composition and longevity of adults were estimated experimentally. The movement intensity was evaluated mostly with the indirect information thus obtained. Females moved more actively than males in all three sawflies.A. japonica females of all age classes moved actively in spring and autumn, but in summer they disappeared. Also,A. rosae females of all age classes moved actively in spring and autumn. In summer, in contrast withA. japonica, A. rosae females moved most actively among the three species in all seasons.A. infumata females, in particular the young females, moved most actively among the three species, exceptA rosae in summer. The movement patterns of the three sawflies were deduced in relation to the spatio-temporal distributions of their habitats. In spring and autumn, when host plants were abundant,A. japonica andA. rosae females were dispersed among the host patches within the census are. In summer, however, when host plants were scarce,A. japonica entered diapause, whereasA. rosae migrated to neighboring areas. On the other hand,A. infumata, in particular young famale, innately dispersed to seek for temporary host plants throughout the census seasons.     

Crucial factors determining the spatio-temporal distribution patterns of threeAthalia sawflies feeding on common cruciferous plants

Summary ThreeAthalia sawflies,A. japonica, A. rosae andA. infumata, feeding on cruciferous plants, coexist in Japan. However, it is not known what ecological strategies they use and what environmental factors are crucial to such strategies. I attempted to explain these questions by examining the relationship between the spatio-temporal distribution patterns of threeAthalia sawflies and their habitats in three districts (Lowland, Intermediate and Mountain). The three sawflies have different spatio-temporal distribution patterns, though they usually used common cruciferous plants.A. japonica was abundant in spring and autumn but disappeared during summer in all the districts. In the Lowland, populations ofA. rosae andA. infumata, like that ofA. japonica, crashed in summer. HoweverA. rosae occurred mainly in summer in the Intermediate and Mountain. AlthoughA. infumata occurred in the same seasons asA. rosae in all districts, population levels ofA. infumata were always lower than those ofA. rosae. The crucial factors controlling their population patterns were the availability of host plants and temperature. Population crashes ofA. rosae andA. infumata were due to food depletion, and those ofA. japonica were due to heat stress. On the other hand, their population patterns may be interpreted as phenological synchronization with their original host plants, though they all existed on common cruciferous plants. The three sawflies may have evolved different strategies to escape from unfavorable habitat conditions. Such strategies are speculated to be summer diapause inA. japonica, long distance migration inA. rosae, and local dispersal inA. infumata.     

Shoot growth dynamics and size-dependent shoot fate of a clonal plant,Festuca rubra, in a mountain grassland

The relation of the within-season and between-season patterns of shoot growth were compared in a clonal grass with long-lived shoots,Festuca rubra, in a mown mountain grassland. The growth rate of shoot length from spring to summer in a year was almost constant for each shoot irrespective of spring shoot length each year. The annual shoot growth rate from spring to spring was negatively correlated with the shoot length in the first spring. Shoots of different length and age therefore tended to converge over time to a population of identical shoot size, suggesting an equalizing effect of growth pattern on size structure. Shoot size (shoot length and number of leaves) influenced the fates of shoots. Larger shoots showed an increased incidence of both flowering and formation of intravaginal daughter shoots and a decreased incidence of death in the subsequent time period. The fates of shoots were independent of their age. Although the negatively size-dependent springto-spring annual shoot growth rate acted to decrease shoot size variation, the remaining variation within the shoot population was still sufficient to generate different fates of shoots. These fates were not related to the previous life history of individual shoots. There was a significantly positive effect of the shoot size at initiation on its life expectancy. This was mainly attributable to the positively size-dependent survival rate of shoots in the early stage (<1 year old) of shoot life history. Later on (> 1 year old), shoot size had little effect on the survival rate of shoots. Once small young shoots have survived this early stage (< 1 year old) in life history, they can grow vigorously, little affected by competition regardless of shoot size, and converge to a stable size structure of shoots of similar size. Only shoot size in the early stage ( < 1 year old) of life history is important for the persistence of a shoot population.     

Host-attacking behavior of a eulophid parasite,Kratochviliana sp., to the leaf mining host,Phytomyza ranunculi (Diptera: Agromyzidae) during its stay on the leaf

Summary The present paper studies how the female parasite ofKratochviliana sp. visits and attacks its host larvae of Ranunculus leaf mining fly,P. ranunculi at a single leaf visit. The parasite visited its hosts at random on the leaf. The frequency of host visits was independent of the host density and the proportion of hosts survived from the parasite attack, in a leaf and its distribution was expressed as a single straight line. It almost always attacked living hosts at the first host visit after isolated from them for one day but with the rate of about 0.5 at the subsequent visits. In consequence, the relationships of the number of host attacks and killed hosts to the host density drew satulated curves in each. A model of host attack by this parasite at its single leaf visit was formulated by modifyingBakker et al.'s model (1972) basing upon these observations and the attack avoidance by the parasite to already attacked hosts previously reported. Ecological studies on the relationship between Ranunculus leaf mining fly,Phytomyza ranunculi Schrank (Diptera; Agromyzidae) and its parasite,Kratochviliana sp. (Hymenoptera; Eulophidae) from the viewpoint of spatial structure II. This paper constitutes a part of the Doctoral Thesis presented to the College of Agriculture, Kyoto University by the present author.     

Mining pattern of the honeysuckle leaf-minerPhytomyza lonicerae

Summary The mining pattern of the honeysuckle leaf-minerPhytomyza lonicerae which feeds onLonicera gracilipes was analyzed with special reference to the avoidance of parasitoids' attacks. The leaf-miner makes the linear mine with branches and crosses by turning and branching, off the backtracking mine. Both average numbers of turnings and branchings were about 4. The speed of mining was 1.5 times faster in backtracking than in advancing. The mine width increased as the mining larva grew and total volume of consumed plant tissue was nearly constant independent of variability of the number of turnings and other characteristics of the mine. The majority of larval mortality were due to attacks by parasitoid wasps (Chrysochairs pentheus andDiglyphus minoeus), which locate the present position of leaf-miners, by tracing mine tracks, and wasps attack leaf-miners at a higher rate in backtracking than in advancing. The adaptive significance of branching and crossing by leaf-miners was discussed in terms of the reduction of parasitoids' attacks.     

Spatial pattern of pine wilt disease caused byBursaphelenchus xylophilus (Nematoda: Aphelenchoididae) within aPinus thunbergii stand

Summary To understand the mechanism of spread of pine wilt disease caused by the pinewood nematode,Bursaphelenchus xylophilus, which is vectored by a cerambycid,Monochamus alternatus, the spatial distribution of trees weakened by the nematode was examined within aPinus thunbergii stand from June to October for 4 years. The weakened trees were distributed in a clumped pattern in 1980 and 1981, at an early stage of infestation. In many cases, they showed a double-clumped pattern. The degree of aggregation was higher in June or July than after August. They were uniformly distributed in June or July 1982 and in June 1983 whereas they showed a double-clumped pattern after August. The trees were frequently weakened in June or July when they were near the trees weakened during the previous year. At quadrat sizes of more than 25 m², spatial overlapping was pronounced between trees weakened during June–July of the current year and those weakened in the previous year. The seasonal changes in spatial distribution of weakened trees were explained by the interaction amongM. alternatus, B. xylophilus andPinus trees.     

Contest and scramble competitions in two bruchid species,Callosobruchus analis andC. Phaseoli (Coleoptera: Bruchidae)

Summary We performed multiple-generation competition experiments betweenCallosobruchus analis andC. phaseoli with different bean sizes. In each system, we supplied 5 g of mung beans (Vigna radiata) every 10 days. We examined three types of bean conditions: 5 g of large beans, 5 g of small beans, and a mixture of 2.5 g of large and small beans. In small bean condition,C. analis dominatedC. phaseoli in all three replicates andC. phaseoli was extinct by the 260th day. On the contrary,C. phaseoli overcameC. analis within 250 days in large beans in all three replicates. In mixed beans condition the two bruchid species coexisted more than 500 days in two out of the three replicates. Even in the exceptional case, both species coexisted for 460 days. These results were examined in the light of the predictions from short-term larval competition experiments and a game theoretical model by Smith and Lessells (1985). The density and frequency dependent results during larval competition inside a bean was concluded to be a main factor to produce the above long-term competition results.     

Prey preference of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand)

Summary Functional responses of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand) attacking the rice brown planthopper,Nilaparvata lugens (St?l.), and the mirid predatorCyrtorhinus lividipennis Reuter were both those of Holling Type II. The attack rate was higher and handling time lower forC. lividipennis. However, when caged with the two prey, the wolf spider showed a significant preference forN. lugens at a lower prey proportion. Proportions of prey attacked were significantly different from the expected ratios of prey available as well as from the predicted preferences derived from the functional response parameters. As proportions ofN. lugens attacked changed from greater to less than expected as the proportions ofN. lugens available increased, a “reverse switch” behaviour seems to be evident.     

Contest competition inDrosophila subobscura

The role of larval intraspecific competition in laboratory populations ofDrosophila subobscura was investigated. Mortality is density-independent during the first 3 days after hatching but becomes density dependent as development proceeds to pupation. Although total biomass per patch was independent of initial egg density, competition betweenDrosophila larvae leads to the formation of smaller pupae. This resulted in a population that was dominated by suppressed individuals. Development rate ofD. subobscura larvae was not affected by high larval densities. Smaller pupae give rise to females with fewer eggs in their ovarioles. A simple simulation model, predicting the effects of intraspecific competition on the fecundity of the nextDrosophila generation is described.     

Binomial sampling plans for estimatingBemisia tabaci populations in cantaloupes

Early season infestations of the sweet potato whitefly,Bemisia tabaci (Gennadius), on cantaloupes,Cucumis melo L., were determined by counts of the number of adults per leaf in fields near Yuma, Arizona. We used these data to develop binomial sampling plans based on the relationship between mean densities of whiteflies per leaf,m, and proportion of leaves infested with more thanI whiteflies,P _I, according to the empirical model lnm=a′+b′ ln[−ln(1−P _I)]. The models were developed for the presence-absence approach (I=0) and for a cutoff value of three whiteflies per leaf (I=3). Four independent data sets were used to evaluate the models. Both methods yielded reliable predictions at low infestation levels, but some of the higherm values were overestimated. As the tentative economic threshold forB. tabaci is three adults per leaf, which corresponds to lowP _I values, results of the binomial sampling were satisfactory for pest management purposes.     

Social structure and the defensive role of soldiers in a eusocial bamboo aphid,Pseudoregma bambucicola (Homoptera: Aphididae): A test of the defence-optimization hypothesis

A complete understanding of the evolution of sociality in aphids requires a detailed knowledge of the patterns of soldier investment in their ecology. The eusocial bamboo aphidPseudoregma bambucicola has a morphologically specialized first-instar soldier caste. The proportion of soldiers was positively correlated with colony size. Within a colony, soldiers were evenly distributed among subcolonies; within each subcolony, however, their distribution was biased toward peripheries which were exposed to many predators. Field experiments introducing natural enemies such asEupeodes confrater (Diptera: Syrphidae) andSynonycha grandis (Coleoptera: Coccinellidae) revealed that the survival rate of these predators was negatively correlated with the density of soldiers, suggesting that soldiers can more or less defend their colonies by killing or removing a range of natural enemies. Observations suggest that large mature colonies attract more predators than newly established small colonies and that, within a colony, the predators attack each subcolony regardless of its position on bamboo shoots. This implies the presence of a positive correlation between colony size and predation risk. Thus, the investment in soldiers seems to reflect the attacking pattern of predators within a colony. These results agree with the defence-optimization hypothesis in soldier investment ofP. bambucicola colonies.     

Host plant conspicuousness and the distribution of eggs and larvae in the butterfly,Anthocharis scolymus (Lepidoptera: Pieridae)

Summary We investigated how the distribution pattern of eggs and larval on the host plant,Turritus glabra, was influenced by the oviposition behavior of the pierid butterflyAnthocharis scolymus. Females searched for the host plants visually and they frequently approached taller host plants with sparse surrounding vegetation. After encountering host plants, oviposition behavior of females was independent of host plant characteristics such as height, density, and type of surrounding vegetation. A female laid eggs singly on a host plants. Most females appeared to lay their eggs regardless of the presense of eggs on the host plant. Consequently egg and larva tended to be abundant on conspicuous host plants as measured by height or relative isolation from other plants. However, overcrowding of eggs on an individual host decreased the survival rate of larvae.     

Accumulation of phyllodulcin in sweet-leaf plants ofHydrangea serrata and its neutrality in the defence against a specialist leafmining herbivore

Among wild plants ofHydrangea serrata (Hydrangeaceae) in Japan, there are sweet plants whose leave contain a kind of isocoumarin, phyllodulcin, which happens to be 350 times as sweet as sucrose to the human tongue. In a primary beech forest in Ashu, Kyoto, the spatial distribution of sweet plants and temporal and the spatial distribution of phyllodulcin within and among plants were investigated using a high performance liquid chromatograph. The distribution of sweet plants was confined within a valley and was parapatric with non-sweet plants. A plant's characteristic phyllodulcin accumulation did not change, even when transplanted into the different habitats. The phyllodulcin content of the sweet plants varied greatly among plants, and the population mean peaked in July when the plants flowered. Within a plant, phyllodulcin content was elevated by partial defoliation. We examined the possible effect of phyllodulcin on herbivory by a specialist leafmining herbivore,Antispila hydrangifoliella (Lepidoptera: Heliozelidae). We transplanted sweet and nonsweet plants reciprocally between their original habitats, excluded attacks by parasitoids, and compared performance of the leafminer. Leafminer colonization and larval survivorship on transplanted andin situ plants was not significantly different between sites. The fact that accumulation of phyllodulcin did not augment a defensive function, at least against herbivory by the leafminer, and the sporadic distribution of phyllodulcin-accumulating plants, suggest that the genotypes synthesizing phyllodulcin emerged independently at separate localities by mutation, and that the genotypes are almost adaptively neutral in defence against the specialist herbivore.     

Comparative studies on the seasonal occurrence, nymphal development and food menu in two giant water bugs,Diplonychus japonicus vuillefroy andDiplonychus major esaki (Hemiptera: Belostomatidae)

Summary We compared the seasonal changes in population density and nymphal development at different water temperatures and under different food conditions between two giant water bugs,Diplonychus japonicus andD. major, in Okayama, Japan. D. japonicus produced 1–2 generations a year, whileD. major was strictly inivoltine. The developmental velocity was higher inD. japonicus than inD. major. The thermal constant ofD. japonicus was less than that ofD. major. These results suggest thatD. japonicus is adapted, to higher water temperature thanD. major. In the field,D. japonicus preyed predominately on Lymnaeidae and Physidae snails, whileD. major preyed on aquatic insects such as dragonfly nymphs.     

Seasonal life cycle and autogeny in the mosquitoAedes togoi in northern Kyushu, Japan, with experimental analysis of the effects of temperature, photoperiod and food on life-history traits

We studied seasonal changes in the larval population structure, adult size, and autogeny (egg production without a bloodmeal) of the mosquitoAedes togoi on the seacoast of northern Kyushu, Japan. The effects of temperature, photoperiod and food conditions on larval development, adult size and autogeny were studied in the laboratory.Aedes togoi overwintered in both the egg and larval stages and was multivoltine. Adult size was greater in spring and autumn and smaller in summer. Autogeny occurred in spring and autumn but not in summer. Autogenous females were larger than anautogenous females, and larger autogenous females produced more eggs. Laboratory experiments showed that autogeny was promoted under low temperature and short-day conditions which corresponded with spring/autumn conditions in the field. Experimental food limitation in the larval stage greatly reduced adult size and autogenous egg production. In the field, large inter-pool variations in adult size and autogeny rate probably resulted from variations in nutrient and crowding conditions. Seasonal autogeny ofAedes togoi was discussed in terms of life-history strategy under fluctuating environmental conditions where relative advantages of autogenous and anautogenous reproduction alternate seasonally.     

Warning coloration in sawflyAthalia rosae larva and concealing coloration in butterflyPieris rapae larva feeding on similar plants evolved through individual selection

Summary Green larvae of the butterflyPieris rapae and black larvae of the sawflyAthalia rosae feed on green leaves of the same cruciferous plants. To demonstrate thatP. rapae has concealing coloration and thatA. rosae has warning coloration, the larvae of the two species were supplied to naive chicksGallus gallus on white, green or black backgrounds.P. rapae larvae were palatable and their green body color acted as a concealing coloration. On the other hand,A. rosae larvae were unpalatable and their black body color acted as a warning coloration. There is a general consensus that warning coloration is an altruistic character which needs victims, and thus can evolve through kin selection or green beard selection. However, blackA. rosae larvae were seldom injured by chicks' attack, in particular, on the green background. Therefore, the warning coloration ofA. rosae larvae can be a selfish character and hence can evolve through individual selection as well as concealing coloration ofP. rapae.     

Evolutionary character changes and population responses in an insect host-parasitoid experimental system

In an insect host (the cowpea weevilCallosobruchus maculatus)- parasitoidHeterospilus prosopidis) experimental system, the population densities of the component species oscillated for the first 20 generations and then abruptly stabilized as the parasitoid density decreased. Examination of the host and parasitoid after the 40th generation in the long-term experiment showed that (1) host larvae exhibited contest-type competition (killing other larvae inhabiting the same bean), in contrast to the founder population being scramble-type competitors and (2) the parasitoid attack rate on the host did not change. There was also an evolutionary trade-off between body size and the rates of larval survival and development, suggesting a cost of contest competition on larval survivorship and development. I tested model predictions (Tuda and Iwasa 1998) that (1) host equilibrium population size should gradually decrease as the proportion of the contest type increases and that (2) random attacks of the parasitoid on the host should reduce the rate of increase in proportion of the contest type, and the effect should become manifest especially during the first 20 generations. Two of three host-only replicates showed significant decrease in population sizes. Although the density of emerging adults per bean did not differ between replicates of the host-only and host-parasitoid systems, comparison of the host body size between them on day 270 (at the 13th generation) showed that the host was more contest-type in the host-only system than in the host-parasitoid system, as the model predicted, and later on day 650 the effect of the parasitoid had disappeared.     

Spatio-temporal distribution patterns of two tachinid flies,Epicampocera succincta andCompsilura concinnata parasitizingPieris

Summary Larvae of genusPieris in the northern part of Kyoto City are parasitized by two tachinid flies:Epicampocera succincta, a specialist on genusPieris, andCompsilura concinnata, a generalist with very wide host-range. We surveyed the parasitism rates ofPieris by both flies for two years at six study areas. In these study areas, there lived three host species in the genusPieris: P. rapae, P. melete, andP. napi, but neither tachinid parasitizedP. napi to any significant extent. In the mountainous district,P. rapae andP. melete coexisted and their populations were relatively continuous, while in the lowland, onlyP. rapae larvae were abundant in spring and autumn, but even they disappeared in summer. Parasitisms byE. succincta occurred mainly in mountainous district and never in the lowland.C. concinnata parasitizedPieris in all the areas, but its parasitisms occurred mainly in autumn. We analyzed the factors affecting the spatial and temporal patterns of parasitism rates and presumed that the temporal discontinuity of host population restricted the distribution of the specialist parasitoid.     

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae) II. Larval competition and interference mechanisms

Summary Competition between contest and scramble strategists was examined using two strains ofCallosobruchus maculatus, the contest strain (iQ) and the scramble strain (tQ). The direct larval interference experiment within each strain showed that the contest strain performed strong direct interference throughout its larval stage against the conspecific opponent(s). The scramble strain also performed its interference, but only during the 3rd and the 4th larval stages. The inter-strain larval competition experiments inside a large and small mung bean (Vigna radiata) showed that the contest strain was, in general, superior to the scramble strain in competition, but the competition results were density- and frequency-dependent. In the large beans, the proportion of adult emergence of the contest strain increased with the increase of its own initial density but decreased as the initial density of the scramble strain increased. The shape of the proportion of adult emergence became nonlinear in the small beans. The cause of the complexity of inter-strain competition results was discussed in the light of the difference in larval interference ability between the two strains.